de Recherche sur l'Environnement Marin
(CEFREM), CNRS/Université de Perpignan,
52 Avenue de Villeneuve, 66860 Perpignan Cedex,
France.
e-mail: ludwig@univ-perp.fr
1. Mackenzie, F. T. & Garrels, R. M. Am. J. Sci. 264, 507–525
(1966).
2. Smith, S. V. & Hollibaugh, G. T. Rev. Geophys. 31, 75–89 (1993).
3. Williams, P. M. & Druffel, E. R. M. Nature 330, 246–248 (1987).
4. Raymond, P. A. & Bauer, J. E. Nature 409, 497–500 (2001).
5. Hedges, J. I., Keil, R. G. & Benner, R. Org. Geochem. 27,
195–212 (1997).
6. Ludwig, W., Probst, J.-L. & Kempe, S. Glob. Biogeochem. Cycles
10, 23–41 (1996).
7. Kao, S. & Liu, K. Limnol. Oceanogr. 41, 1749–1757 (1996).
8. McClain, M. E., Richey, J. E. & Brandes, J. A. Glob. Biogeochem.
Cycles 11, 295–311 (1997).
number of legs, humans and quadrupeds
change from walk to run or trot at the same
Froude number, close to 0.5 (ref. 3). Even
gibbons
8
change from merely swinging to
swinging with aerial phases at Froude num-
bers ranging from 0.3 to 0.6. Humans of
short height, such as children
9,10
, patients
suffering from early-onset growth-hormone
deficiency
11
and pygmies
12
, optimally walk
at speeds that correspond dynamically to
that for adults, that is, with a Froude number
of 0.25. Here, walking speed is optimal when
the recovery of energy by the body — by
exchanging potential and kinetic energies —
is maximal
7
. What all of this means is that,
within the same gravitational environment
(such as on the Earth), the smaller the body,
the lower the ‘equivalent’ speed of move-
ment, which is proportional to the square
root of the leg length.
But the power of the Froude number for
predicting equivalent walking speeds is
not confined to the Earth. Within the same
species and for a given body size, dynamic
news and views
Presynaptic active zones are
the specialized sites from
which nerves release
neurotransmitter. They are
characterized by the presence
of calcium channels, synaptic
vesicles containing
neurotransmitter, and
aggregates of protein that
make up what is known as
‘active-zone material’. This
material was described by
S. L. Palay almost 50 years ago
(J. Biophys. Biochem. Cytol. 2,
193–202; 1956), but its precise
structure and function have
remained an enigma. Things
may be about to change,
however, thanks to the
publication elsewhere in this
issue of stunning three-
dimensional reconstructions of
the frog neuromuscular junction
(Nature 409, 479–484; 2001).
To study this junction
between nerve and muscle,
M. L. Harlow and colleagues
used a technique known as
electron microscope
tomography, which gives
greater spatial resolution than
conventional transmission
electron microscopy. They
describe three distinct elongate
structures — which they call
ribs, beams and pegs — that
make up the first 15 nanometres
of active-zone material as one
looks into the neuron. The top
image shows this view, with
the active-zone material in
yellow and the synaptic
vesicles in blue.
Beams — which make up
the vertical band in the centre
of this image — are arranged
along the long axis of the
elongate active zone, and form
the backbone of the structure.
Ribs lie perpendicular to this
long axis, and link the central
backbone to the double row of
synaptic vesicles that straddle
it. Pegs (not apparent in these
images) tether each rib to the
plasma membrane in a pattern
that matches the distribution of
molecules that are thought to
be calcium channels. The lower
image is a side-on view, with
the presynaptic plasma
membrane in grey. The
structure is perhaps best
appreciated in the movies
published on Nature’s web site
(http://www.nature.com/nature/).
As a whole, the active-zone
material appears to provide a
scaffold by which synaptic
vesicles are localized to the
specialized presynaptic
membrane. But the proteins
that make up the structure
probably have more specific
functions. The intracellular
domains of calcium channels
may form all or part of the pegs,
and ribs may be composed of
the host of proteins that
together mediate docking and
fusion of vesicles with the
presynaptic membrane. We
eagerly await the formal
identification of the rib, beam
and peg proteins. Lesley Anson
Neurobiology
Activity at the active zone
Biomechanics
Walking on other planets
Alberto E. Minetti
A nineteenth-century equation used for building model ships allows us to
compare the motion of animals of different sizes and gaits. It may also
give us an idea of how we would move on different planets.
W
hat do a gibbon swinging by its arms
from tree to tree, a sailing steamship,
and a walking human have in com-
mon? The answer lies in a simple concept
introduced in the nineteenth century by
nautical engineer William Froude to help
him to produce model ships that maintained
the same propulsion dynamics as full-size
vessels. His ‘Froude number’ is a dimension-
less variable that was brought into biology
by D’Arcy Thompson
1
and popularized by
McNeill Alexander
2,3
in the study of the
energetics and mechanics of animal loco-
motion. This number allows one to compare
the motion of species with different numbers
of legs and gaits, and to investigate the effects
of different body sizes on the mechanics of
movement. As suggested by various earlier
papers
4–6
, and borne out by new work
described by Cavagna and colleagues in the
Journal of Physiology
7
, the Froude number
is also a reliable rule-of-thumb for predict-
ing walking speeds on planets with gravities
different from that of the Earth.
In walking humans or swinging gib-
bons, changes in the vertical position of the
body’s centre of mass during movement
affect the gravitational potential energy of
the body, and are accompanied by opposite
changes in the kinetic energy needed to
drive movement. The result is a pendulum-
like mode of movement that saves mech-
anical energy. Roughly the same principles
apply to ships, but here the potential energy
is related to the size of the wave generated
by the ship.
One of the theories underlying studies
of movement — dynamic similarity
3
—
basically states that geometrically similar
bodies that rely on pendulum-like mech-
anics of movement will have similar gait
dynamics if the Froude number remains
the same. This number (Fr) is given by
Frǃv
2
/(gǂl), where v is the speed of move-
ment (in m s
ǁ1
), g is acceleration due to
gravity (in m s
ǁ2
) and l is a characteristic
length (such as leg length, in metres). The
Froude number is directly proportional to
the ratio between the kinetic energy and
the gravitational potential energy needed
during movement.
Dynamic similarity implies that, for
example, despite differences in body size and
NATURE | VOL 409 | 25 JANUARY 2001 | www.nature.com 467
M. L. HARLOW ET AL.
© 2001 Macmillan Magazines Ltd